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Aplexa Hypnorum (Gastropoda: Physidae) Exerts Competition on Two Lymnaeid Species in Periodically Dried Ditches

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Aplexa Hypnorum (Gastropoda: Physidae) Exerts Competition on Two Lymnaeid Species in Periodically Dried Ditches Ann. Limnol. - Int. J. Lim. 52 (2016) 379–386 Available online at: Ó The authors, 2016 www.limnology-journal.org DOI: 10.1051/limn/2016022 Aplexa hypnorum (Gastropoda: Physidae) exerts competition on two lymnaeid species in periodically dried ditches Daniel Rondelaud, Philippe Vignoles and Gilles Dreyfuss* Laboratory of Parasitology, Faculty of Pharmacy, 87025 Limoges Cedex, France Received 26 November 2014; Accepted 2 September 2016 Abstract – Samples of adult Aplexa hypnorum were experimentally introduced into periodically dried ditches colonized by Galba truncatula or Omphiscola glabra to monitor the distribution and density of these snail species from 2002 to 2008, and to compare these values with those noted in control sites only frequented by either lymnaeid. The introduction of A. hypnorum into each ditch was followed by the progressive coloni- zation of the entire habitat by the physid and progressive reduction of the portion occupied by the lymnaeid towards the upstream extremity of the ditch. Moreover, the size of the lymnaeid population decreased significantly over the 7-year period, with values noted in 2008 that were significantly lower than those recorded in 2002. In contrast, the mean densities were relatively stable in the sites only occupied by G. truncatula or O. glabra. Laboratory investigations were also carried out by placing juvenile, intermediate or adult physids in aquaria in the presence of juvenile, intermediate or adult G. truncatula (or O. glabra) for 30 days. The life stage of A. hypnorum had a significant influence on the survival of each lymnaeid. In snail combinations, this survival was significantly lower for adult G. truncatula (or O. glabra) than for intermediate snails. In contrast, the survival of juveniles was similar to that noted in the corresponding controls. This interspecific competition between the physid and either lymnaeid may not be due to food present in their habitat, but might be due to the toxicity of mucus secreted by intermediate and adult A. hypnorum. Key words: Aplexa hypnorum / colonization / density / ditch / Galba truncatula / Gastropoda / Lymnaeidae / Omphiscola glabra / Physidae Introduction populations because of the summer drought which was too short: 5–6 weeks (Rondelaud et al., 2006). A terrestrial Fasciolosis due to Fasciola hepatica (Linnaeus) has a snail, Zonitoides nitidus (O.F. Mu¨ ller), able to actively worldwide distribution, with human and animal cases prey on these lymnaeids, was also applied in several cattle- being reported on the five continents. To accomplish its breeding farms. However, this method has not become life-cycle, this fluke needs a definitive host (generally a generalized because of the complexity of applying this mammal) that harbours the adult parasite and an inter- control in the field by non-specialists (Rondelaud et al., mediate host (a freshwater pulmonate gastropod) that 2006). ensures the development of larvae. In Western Europe, the Another method of control is to use a freshwater snail, most common host snail is Galba truncatula (O.F. Mu¨ ller) which is not a compatible host for F. hepatica,asa (Taylor, 1965; Torgerson and Claxton, 1999). Another competitor of these lymnaeid populations. Several gastro- lymnaeid, Omphiscola glabra (O.F. Mu¨ ller) also can sus- pods have already been used as competitors in the past tain larval development of F. hepatica (Abrous et al., 1999, to control intermediate hosts for schistosomes, but the 2000; Rondelaud et al., 2015). The control of these success of these biological agents varied according to the invertebrate populations has been proposed by numerous snail species and the country in which they were intro- authors as one of the means to eliminate the fluke from duced (Brown, 1994). As no freshwater pulmonate gastro- farms (Taylor, 1965). On the acidic soils of central France, pod was reported as a competitor of G. truncatula and the use of open drainage only limited the size of snail O. glabra in the literature, it was reasonable to search for a candidate among species belonging to the same *Corresponding author: [email protected] snail community. In Western Europe, these lymnaeids are Article published by EDP Sciences This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited 380 D. Rondelaud et al.: Ann. Limnol. - Int. J. Lim. 52 (2016) 379–386 known to live in periodically dried habitats with the physid Aplexa hypnorum (Linnaeus) and planorbids such as Anisus leucostoma (Miller) or A. spirorbis (Linnaeus) (Costil et al., 2001; Vignoles et al., 2015). As another physid, Physella acuta (Draparnaud) was already reported as a competitor against Bulinus truncatus (Audouin) in Egypt (El-Hassan, 1974), the competition of A. hypnorum against G. truncatula and O. glabra was investigated. A. hypnorum is known for its distribution in most European countries (Welter-Schultes, 2012, 2013) and North America (Dillon et al., 2006). But in some countries, the numbers of these snails were small such as in woodland ponds from Southern Poland (Spyra, 2010) and a few ponds of the upper Rhine valley (Mosimann, 2000; Bauer and Ringeis, 2002). This physid colonizes small and temporary waterbodies rich in veg- etation, located in lowlands. It is also found on swampy and forested margins of lakes, in swampy deciduous forests, and very moist meadows (Welter-Schultes, 2013). However, the species prefers ditches, temporary ponds Fig. 1. Location of the ten road ditches in the Brenne Regional and pools, which it shares with other gastropod species Natural Park, department of Indre (Central France). adapted to seasonal habitats (Kerney, 1999; Anderson, 2006). The bottom of these habitats often consisted of Materials and methods mud (Glo¨ er and Diercking, 2010). In addition, a positive relationship was noted in Germany between the abun- Snail identification was done on the basis of shell shape dance of snails and sand and/or clay soil types (Den and size, and also length of reproductive organs according Hartog, 1963; Glo¨ er and Diercking, 2010). The future of to the keys by Glo¨ er and Meier-Brook (2003). A. hypnorum is particularly worrying in Western European countries due to destruction of its moist habitats owing to land drainage and the general intensification of agriculture Field investigations (Kerney, 1999). In these periodically dried habitats, community dy- Snail habitats namics depend on species-specific abilities to withstand drought periods (Dillon, 2004). However, periodic The field part of the investigation included a total of drought is not the sole factor that acts on the dynamics ten ditches inhabited by either G. truncatula or O. glabra. of these populations, as the composition of this freshwater These ditches were located within the municipalities pulmonate community also has also an effect on variations of Chitray, Luzeret, Migne´ and Thenay, department of in snail numbers. In the Brenne Regional Natural Park Indre, Central France (Fig. 1). The first six ditches (Central France), the densities of adult O. glabra were were only inhabited by G. truncatula in March 2002. At similar when this lymnaeid was living alone or with this date, 50 adults (shell height >12 mm) of A. hypnorum A. spirorbis in its habitat. In contrast, the size of each from a nearby upstream pond (46x45k14kkN, 1x11k7kkE) O. glabra population fell strongly when A. hypnorum or were introduced in the deepest zone of each of three P. acuta occupied the same habitats (Vignoles et al., 2015). G. truncatula ditches (46x35k53kkN, 1x27k27kkE; 46x36k38kkN, In view of these results, the two following questions 1x27k10kkE; 46x40k33kkN, 1x22k44kkE). The remaining arose: was competition for trophic resources between three G. truncatula-only ditches (46x35k2kkN, 1x27k18kkE; A. hypnorum and O. glabra the reason for the observed 46x36k40kkN, 1x27k8kkE; 46x40k27kkN, 1x21k23kkE) served as population dynamics in coexistence? Might the numerical controls. The length and area of these first six habitats decrease of O. glabra result from a lethal action of ranged from 42 to 87 m and from 18.9 to 39.1 m2, respec- A. hypnorum on eggs and juveniles of the lymnaeid? tively, in March 2002. Following the same methods To answer the first question, field investigations were as above, 50 adult A. hypnorum were introduced to carried out during seven successive years to monitor the the deepest zone of each of two O. glabra-only ditches dynamics of G. truncatula and O. glabra populations living (46x33k37kkN, 1x24k18kkE; 46x40k17kkN, 1x18x3kkE), while in ditches where A. hypnorum was experimentally intro- the other two O. glabra ditches (46x34k47kkN, 1x24k34kkE; duced. The answer to the second question was provided 46x42k47kkN, 1x19k69kkE) served as controls. The length by monitoring population dynamics in laboratory aquaria of these last four habitats ranged from 81 to 146 m in with A. hypnorum and G. truncatula or O. glabra. In March 2002 and their area from 36.4 to 65.7 m2. The both cases, controls were road ditches colonized by longest distance between these ten snail habitats was G. truncatula or O. glabra (but without A. hypnorum), or 18 km. All these ditches contained rain water during most aquaria containing G. truncatula or O. glabra only. of the year (from the end of October to mid-June). D. Rondelaud et al.: Ann. Limnol. - Int. J. Lim. 52 (2016) 379–386 381 The bottom sediment was composed of silt and sand, Kruskal–Wallis test was used to compare snail densities and was supported by impermeable subsoil generally noted in each quadrant in 2002 and 2008. All the statistical formed by sandstone and/or clay. Water pH varied from analyses were performed using Statview 5.0 software (SAS 6.7 to 7.8 and the dissolved calcium level ranged from 26 Institute Inc., Cary, NC, USA). to 35 mg.Lx1 (Dreyfuss et al., 2010).
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